Male infanticide and paternity analyses in a socially natural herd of Przewalski's horses: Sexual selection?

The sexual selection hypothesis explains infanticide by males in many mammals. In our 11-year study, we investigated this hypothesis in a herd of Przewalski's horses where we had witnessed infanticidal attacks. Infanticide was highly conditional and not simply linked to takeovers. Attacks occurred in only five of 39 cases following a takeover, and DNA paternity revealed that, although infanticidal stallions were not the genetic fathers in four cases out of five, stallions present at birth did not significantly attempt to kill unrelated foals. Infanticide did not reduce birth intervals; only in one case out of five was the infanticidal stallion, the father of the next foal; mothers whose foals were attacked subsequently avoided associating with infanticidal stallions. Therefore, evidence for the sexual selection hypothesis was weak. The “human disturbance” hypothesis received some support, as only zoo bred stallions which grew up in unnatural social groups attacked foals of mares which were pregnant during takeovers.     

半散放条件下普通斑马的种群特征

<正>普通斑马又称平原斑马(Equus burchelli),隶属于奇蹄目马科,是一种极具观赏价值的大型哺乳动物。广泛分布于非洲,喜欢栖息在平原和草原,随季节和食物迁徙。普通斑马是社会性动物,习惯群体生活,规模较小的群体由1只雄性和2-6只雌性以及它们未成年的子女组成;或是由十几只未成年的雄性普通斑马组成;较大的群体则有几百只,它们通常是由若干个小规模的群体组成。雄性     

A case of adoption and allonursing in captive plains zebra (Equus burchellii)

Although allonursing (allowing non-filial offspring to suckle) can be a costly behaviour, it has been reported for many mammals including ungulate species. However, such behaviour is very rare in equids. This is the first report on adoption and allonursing in captive plains zebra (Equus burchellii), recorded in the Dv?r Králové Zoo, Czech Republic. We observed a case of adoption of an orphaned foal by a lactating mare, who then regularly nursed two foals (filial and non-filial). The allonursing mare rejected more suckling attempts, terminated suckling bouts more often, and had a shorter suckling bout duration than other mares. When nursing both foals at the same time, the suckling bout lasted for less time than when nursing a single foal, regardless of whether it was filial or non-filial. The allonursing mare apparently did not discriminate between the filial and non-filial foal.     

Maternal care and foal social relationships in a herd of Sorraia horses: Influence of maternal rank and experience

The influence of maternal rank and experience on patterns of maternal care and social relationships of foals were investigated in a managed herd of Sorraia horses, Equus caballus. Social interactions and spatial relationships of 13 foals (seven females and six males) born to seven mares were examined from birth to 10 months of life, within the three major periods of foal development. Conflict over suckling between dam and foal was not generally affected by rank and experience, but higher-ranking mothers allowed more suckling during late lactation than lower-ranking mothers. Foals of higher-ranking mares spent more time in proximity to the mother during socialization. Maternal rank and experience did not significantly affect maternal protectiveness, foal independence from the mother or the development of affiliative relationships between foals and group members. Foals of higher-ranking mares received lower frequencies of aggression from other horses only in the first month of life. Dominance relationships among foals depended mainly on aggressiveness and were not associated with maternal rank. The large variability in maternal behaviour, the absence of a significant association between maternal rank and body condition at parturition and the stable social environment within this herd may partly account for the reported results.     

Further evidence for sex differences in suckling behaviour of captive plains zebra foals

Sex differences in suckling behaviour could be explained either by different maternal investment or by different behaviour of young. Previously, we found sex differences in rejection and termination of suckling bouts in captive plains zebras (Equus burchellii). In this study, we analysed the length of suckling bout interruptions and unsuccessful subsequent attempts which followed a successful suckling bout in plains zebra foals at Dvůr Králové Zoo. When a suckling bout was interrupted, male foals resumed the bout sooner than female foals. The probability of a subsequent attempt decreased with increasing age in female foals but did not change in male foals. These results demonstrate higher effort in suckling interest by male foals, who tried to suckle more than female foals. Although we found sex bias in the effort to suckle, which suggested higher parent–offspring conflict in male offspring, we cannot omit a sex bias in milk supply by the mother.     

Sex preferences in Jeju pony foals (Equus caballus) for mutual grooming and play-fighting behaviors

We investigated mutual grooming by Jeju pony (Equus caballus) foals to determine whether male foals preferentially interact with potential future sexual partners or competitors. We predicted that relative to female foals, male foals would exchange grooming more frequently with young mares and that in general, foals would mutually groom more frequently with the opposite sex rather than the same sex. Observing 53 foals between April and October 1998, we recorded 113 mutual grooming events. Male foals exchanged grooming with yearling mares more frequently than with their mother, while female foals exchanged grooming with their mother more frequently than with yearling mares. Contrary to the prediction, foals were not more likely to mutually groom with a foal of the opposite sex than with a foal of the same sex. In our study, 21 instances of play-fighting behavior followed mutual grooming between peers. Relative to intersexual grooming events, play-fighting was more likely to follow intrasexual mutual grooming, and male foals were much more likely to play fight than female foals. These results provide evidence that Jeju pony foals develop and maintain social relationships at the earliest stage of their lives. We suggest that early social experiences might influence social bonding later when the male foal begins to form a harem after separation from its mother.     

Reproductive performance in Ohio dairy herds in the 1990s

The objective of this study was to evaluate trends in reproductive performance in Ohio dairy herds during the 1990s. Stringent criteria for herd inclusion were applied to DHI records to ensure high quality and reliability of the data. The final data set contained 44,425 monthly herd summary records from 1772 herds from years 1992, 1994, 1996 and 1998. Outcomes of interest were calving interval length, calving-to-conception interval, days to first breeding, services per conception and first service conception rate. PROC MIXED in SAS (version 8.1) was used and the monthly records were treated as repeated measures nested within years and herds. First-order autoregressive covariance structure was used to model the covariance between the repeated measures within herds. The data were analyzed for the entire herd as well as for the first lactation cows separately. During the 1990s reproductive performance declined in Ohio dairy herds. Herd average calving interval lengthened from 13.6 to 14.1 months and calving-to-conception interval increased from 136 to 150 days between 1992 and 1998, respectively. Days to first service and number of services increased while first service conception rate declined on a herd level. The reproductive performance of first lactation cows, however, remained constant during this same period. Overall herd reproductive performance was associated with herd size and fat-corrected rolling herd average milk production (FCM). As milk yield increased, first service conception rate decreased and services per conception increased. Larger herds had lower first service conception rate and needed more services per conception, but they started breeding their cows earlier resulting in a slightly shorter calving interval and calving-to-conception interval compared to smaller herds.     

Birth sex ratios relate to mare condition at conception in Kaimanawa horses

Several hypotheses have been proposed to explain variation inbirth sex ratios, based on the premise that variation is expectedwhen the profitability of raising sons and daughters variesbetween individual parents. We tested the Trivers-Willard hypothesisthat mothers in better condition produce relatively more sonsand that mothers in poorer condition produce relatively more daughterswhen male reproductive success is more variable. We examinedbirth sex ratios in relation to mare body condition at conceptionin horses in which male reproductive success is differentiallyhelped by slight advantages in condition. Horses meet the assumptionsof the Trivers-Willard hypothesis better than many species onwhich it has been tested and in which sex ratio biases are notconfounded by sexual size dimorphism such that one sex is more likelyto die in utero in females in poor condition. Mares that hada female foal were in poorer condition at conception than thosethat had a male foal, and mares that had foals of differentsexes in different years were in significantly poorer conditionwhen they conceived their female foal. There was no relationshipbetween offspring sex and mid-gestation condition, and therewas no difference in foaling rates in relation to body conditionat conception. Consequently, sex ratio deviations are not explainedby fetal loss in utero. Furthermore, differential fetal lossof the less viable sex cannot explain the greater proportionof males produced by mares in better condition. Therefore, ourresults suggest that sex ratio modification occurs at conceptionin wild horses.     

Urinary androgen concentrations and social status in two species of free ranging zebra (Equus burchelli and E. grevyi)

In both species of zebra, breeding males had higher urinary androgen concentrations (ng androgens/mg Cr) than did non-breeding bachelor males (30.0 +/- 5.0 (N = 9) versus 11.4 +/- 2.8, (N = 7) in the plains zebra; 19.0 +/- 2.2 (N = 17) versus 10.7 +/- 1.2 (N = 14) in the Grevy's zebra). In the more stable family structure of the plains zebra (single male non-territorial groups) variations in androgen concentrations could not be ascribed to any measured variable. In the Grevy's zebra, androgen values were significantly lower in samples taken from territorial (breeding) males which had temporarily abandoned their territories (N = 4) and the urinary androgen concentration for a male on his territory was negatively correlated with the time since females last visited the territory.     

Pedigree and herd characterization of a donkey breed vulnerable to extinction

Most donkey and local horse breeds are vulnerable to extinction as mechanization of agriculture progress throughout the world. The present study analyzed the pedigree and herd records of the donkey Asinina de Miranda breed (RAM), identifying genealogical and human factors that may affect the breed genetic diversity in the future and suggesting suitable strategies to breed preservation, early on the conservation program. The breeding rate was very low, with a ratio of foaling/live animals of 0.23 (178/760). The estimated number of founders and ancestors contributing to the reference population was 128 and 121. The number of founder herds in the reference population was 64, with an effective number of founder herds for the reference population of 7.6. The mean age of herd owners was 65.50±0.884 years, with a negative association among the herd size and owner’s age (P<0.001). In contrast, the size of the herd and the ownership of a male were both positively associated (P<0.001) with the herd number of in-born foals. Both the owners’ age and the herd location (RAM home region v. dispersal region) were negatively associated with the foaling number (P<0.001). The main identified risk factors were: low breeding rates; low number of males and their unequal contribution to the genetic pool; unequal contribution of the herds to genetic pool; and advanced age of herd owners.     

Infanticide in <Emphasis Type="Italic">Cercopithecus mitis stuhlmanni</Emphasis> in the Kakamega Forest,Kenya: Variation in the Occurrence of an Adaptive Behavior

Infanticide by males is widespread across mammals and especially prevalent among primates. Considerable research has examined how fitness benefits can explain the occurrence of this behavior; less is known, however, about intrapopulation variation in its occurrence. We evaluated 10 infanticides by males in wild blue monkeys according to the sexual selection hypothesis. To explore intrapopulation variation in occurrence of infanticide, we compared these cases to 38 cases that were contextually similar but in which infanticide did not occur. We examined male reproductive benefit, infant age, maternal parity, postconception estrus, group defense, available mating partners, and context of takeover. We based comparisons on daily or near daily records of male presence in the study groups, infant birth dates, and male-female sexual interactions. Infanticides followed predictions of the sexual selection hypothesis: males were unlikely to kill their own offspring, the period for the mother’s return to conception was reduced by half, and males increased their chance of siring her next offspring. Difference in male reproductive benefit, costs, and motivation did not fully explain the observed variation in infanticide occurrence. Infants were more likely to be spared if they were older when a male first arrived, or if their mother had mated with the male in the second month after conception. The most important determinant of infant fate, however, was male identity, a finding consistent with 2 scenarios: 1) an infanticidal tendency may be influenced by a genetic polymorphism that is not fixed in this population or 2) infanticidal behavior may be a conditional male strategy. Further research on intrapopulation variation in infanticidal behavior should focus especially on characteristics of males.     

Aggression and social spacing in light horse (Equus caballus) mares and foals

Aggression and social spacing were studied in 14 light horse mares and their foals living at pasture. Focal samples were collected on each mare-foal dyad for 6 to 10.5 h from 2 months of foal age until weaning at approximately 4 months of age. Observations on foals continued until approximately 6 months of age for 7.5 to 10.5 h per foal. Every 2 min the identities of all individuals within 5 m were recorded. All occurrences of agonistic behavior, and the participants, were recorded during the focal samples. In addition, during feeding of supplemental grain, all occurrences of agonistic behavior by all subjects were recorded. Significant correlations were found between mare rank and the rank of foals both prior to and after weaning. Before weaning, the rank of the foal was significantly correlated with birth order. No significant correlation between birth order and foal rank was found for the post-weaning hierarchy. An animal's gender had no significant effect on foal rank or the choice of preferred associate. Both prior to and after weaning, foals associated preferentially with the foal of their dam's most preferred associate. In addition, significant positive correlations were found between rank of mares and foals and the rate at which they directed aggression to other herd members.     

New records of within-group infanticide and cannibalism in wild chimpanzees

Two cases of within-group infanticide and cannibalism were observed among the M Group chimpanzees of the Mahale Mountains, Tanzania. In both cases, victimized infants were male, 5 – 6 months of age, and in good health when killed. Four to five years have passed since the mothers of the victims immigrated into M Group as nulliparous immigrants. In one case the 2nd-ranking male was observed to detach the infant from the mother's belly. Both infants were finally killed by the alpha male after several adult males scrambled for the bodies. There was no evidence that the mothers had mated with males other than those of M Group. Nor was there evidence that the mothers had restrictive mating relationships with some of the M Group adult males. What little evidence is available shows that the mothers had mated mostly with adolescent and other immature males during their conception cycles. However, at least in one case, the mother began to mate more with adult males rather than with immature males after the infanticide. It is proposed that the function of within-group male infanticide can be explained by the male-male competition hypothesis developed for hanuman langurs and other nonhuman primates.     

Distinct preoptic‐BST nuclei dissociate paternal and infanticidal behavior in mice

Paternal behavior is not innate but arises through social experience. After mating and becoming fathers, male mice change their behavior toward pups from infanticide to paternal care. However, the precise brain areas and circuit mechanisms connecting these social behaviors are largely unknown. Here we demonstrated that the c‐Fos expression pattern in the four nuclei of the preoptic‐bed nuclei of stria terminalis (BST) region could robustly discriminate five kinds of previous social behavior of male mice (parenting, infanticide, mating, inter‐male aggression, solitary control). Specifically, neuronal activation in the central part of the medial preoptic area (cMPOA) and rhomboid nucleus of the BST (BSTrh) retroactively detected paternal and infanticidal motivation with more than 95% accuracy. Moreover, cMPOA lesions switched behavior in fathers from paternal to infanticidal, while BSTrh lesions inhibited infanticide in virgin males. The projections from cMPOA to BSTrh were largely GABAergic. Optogenetic or pharmacogenetic activation of cMPOA attenuated infanticide in virgin males. Taken together, this study identifies the preoptic‐BST nuclei underlying social motivations in male mice and reveals unexpected complexity in the circuit connecting these nuclei.     

Female resistance to invading males increases infanticide in langurs

Background

Infanticide by adult male occurs in many mammalian species under natural conditions, and it is often assumed to be a goal-directed action and explained predominately by sexual selection. Motivation of this behavior in mammals is limitedly involved.

Methodology and Principal Findings

We used long-term reproductive records and direct observation in captivity and in the field of two snub-nosed langur species on the basis of individual identification to investigate how infanticide happened and to be avoided in nonhuman primates. Our observations suggested that infanticide by invading males might be more accidental than goal-directed. The invading male seemed to monopolize all the females including lactating mothers during takeovers. Multiparous mothers who accepted the invading male shortly after takeovers avoided infanticide in most cases. Our results conjectured primiparous mothers would decrease infanticidal possibility if they sexually accepted the invading male during or immediately after takeovers. In the studied langur species, voluntary abortion or mating with the invading male was evidently adopted by females to limit or avoid infanticide by takeover males.

Conclusions and Significance

The objective of the invading male was to monopolize all adult females after his takeover. It appeared that the mother''s resistance to accepting the new male as a mating partner was the primary incentive for infanticide. Motivation analysis might be helpful to further understand why infanticide occurs in primate species.     

Infanticide risk and infant defence in multi-male free-ranging sooty mangabeys, Cercocebus atys

For years, infanticide by males was thought to be unlikely in multi-male primate species. Recent studies have, however, presented evidence of infanticide in such species and a recent model by Broom and colleagues predicts that males’ age and rank influence the occurrence of infanticide: youngest and highest-ranking immigrant males are more likely to commit infanticide than their older and lower-ranking counterparts if putative fathers fail to protect infants. I collected data on adult free-ranging sooty mangabey females in the Taï National Park, Ivory Coast, over 11 months including a birth and a mating season. Infanticide had been previously reported in captivity for this species, but not in the wild. Several males entered the group prior to and during the mating season. As predicted by the model, only the more dominant immigrant ones attacked mother-infant pairs significantly more often than did other males. Mothers often reacted with counter-attacks. Potential fathers guarded and supported infants and mothers throughout the period of infant vulnerability. Furthermore, as only one of seven infants died despite 136 observed attacks on mother-infant pairs and unattended infants by immigrant males, we conclude that cooperation between putative fathers and mothers represents an effective protection against infanticide.     

Evaluation of a contract breeding management program in Ohio dairy herds: test day summary and economic measures

A field study was conducted to assess the impact of a contract breeding program that was offered by a breeding co-operative and featured tail chalking and daily evaluation of cows for insemination by co-operative technicians; dairy employees no longer handled estrous detection and insemination activities. From early 2002 until mid-2004, herd-level test day summary records related to production and reproduction were obtained for 32 herds identified as well-managed client herds of the breeding co-operative. Using analyses that controlled for other predictors and random herd-level effects, average days to first service were less by 13 days (P=0.0037) and estrous detection rate was greater by 12% (P=0.0011) for program than for non-program herds. Although first service conception rate was slightly less and the program herds used 0.34 more services per conception (P=0.1488) than non-program herds, the program herds averaged 16 fewer days before pregnancy (P=0.028). Test day summary information and representative estimates of feed, milk, and semen prices were used in a spreadsheet-based model to estimate a partial budget annuity value for an average cow in each herd on each test day. Value of an average cow from a contract herd did not significantly differ from a non-contract herd, even though the analyses suggested an economic benefit for the program herds; the modeling did not, however, account for costs of the program implementation. Additional analyses did not find any significant associations between technician and on days to first service, first service conception rate, estrous detection rate, services per conception, or days open.     

Infanticide among free-ranging langurs (Presbytis entellus) at jodhpur (Rajasthan/India): Recent observations and a reconsideration of hypotheses

Considerable disagreement characterizes the debate concerning frequency, causation, and function of infanticide in connection with adult male replacements in bisexual one-male troops of hanuman langurs (Presbytis entellus). Detailed observations are presented about two noninfanticidal and three infanticidal male changes including six eye-witness and five presumed cases of infanticide within three langur troops during a long-term study at Jodhpur, Rajasthan, India. The results do not support any explanatory hypotheses focussing on social crowding, regulation of population density, social stress, sexual frustration, incest avoidance, or social bonding, but are in general though not total agreement with the reproductive advantage hypothesis: mainly unrelated infants were killed (one possible exception), the infanticidal male generally sired the subsequent offspring (one exception), and the mean interbirth interval subsequent to infanticide is by 2.1 months shortened. Likewise, several cases of stress induced abortions occurred. It is demonstrated that postconception estrous behaviour is by no means a female counterstrategy to infanticide in order to confuse males concerning the issue of paternity, since an infanticidal male did not spare the subsequent offspring of mothers who copulated with him during pregnancy and pregnant females did not discriminate between fathers and non-fathers.     

Dynamic spatial partitioning and coexistence among tall grass grazers in an African savanna

Competitive relationships among mobile animals may be expressed through dynamically changing spatial relationships over different time frames. Less common species that are apparently inferior competitors may be able to coexist with more abundant species by concentrating in regions of the landscape little utilized by the former at spatio‐temporal scales from annual or seasonal ranges to the specific foraging localities exploited at different stages of the annual cycle. Spatial relationships may be influenced further by dependencies on other resources, predation risks and facilitatory interactions under certain conditions. Our study aimed to determine whether competition with more abundant zebra and buffalo restricted the abundance of sable antelope in a region where these three tall‐grass grazers overlapped in their herd distributions. We tracked the simultaneous movements of animals representing herds of these species over two dry seasons and one wet season using GPS‐GSM collars, and estimated seasonal or monthly range extents and their overlap. We also compared daily separation distances between these animals against the null pattern expected if their movements had been independent, and assessed how prior grazing by buffalo influenced the subsequent use of these localities by sable. The range of the sable herd was mostly separated from the seasonal range of the buffalo herd during the late dry season of 2006 and throughout the dry season of 2007. Seasonal home ranges of zebra herds overlapped partially with the range of the sable herd during most of the year. Even during times when their ranges overlapped, sable were rarely recorded within <1 km of the buffalo herd. Prior grazing by buffalo beyond a threshold level inhibited later use of these localities by sable, but the sable were nevertheless able to exploit places that were little utilized by buffalo at that time. Sable were less able to evade overlap with the small, mobile zebra herds, and hence more vulnerable to competitive exclusion by zebra than by buffalo. Our findings demonstrate how less abundant species can restrict competition from more abundant competitors through dynamic spatial partitioning in regions where their home ranges overlap.     

A rare fight in female plains zebra

We describe a fight between two female plains zebra (Equus burchelli). Plains zebra are ungulates with stable social groups known as harems. Female aggression rarely escalates to the level we observed. The fight immediately followed the birth of a foal to one of the females. The initiating female repeatedly kicked and bit the mother, who reacted aggressively and by guarding her foal. We present hypotheses on the causes underlying this rare event.